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Molecular Panbiogeography of the Tropics

Molecular Panbiogeography of the Tropics

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Molecular Panbiogeography of the Tropics

902 pages
11 heures
Jan 4, 2012


Molecular studies reveal highly ordered geographic patterns in plant and animal distributions. The tropics illustrate these patterns of community immobilism leading to allopatric differentiation, as well as other patterns of mobilism, range expansion, and overlap of taxa. Integrating Earth history and biogeography, Molecular Panbiogeography of the Tropics is an alternative view of distributional history in which groups are older than suggested by fossils and fossil-calibrated molecular clocks. The author discusses possible causes for the endemism of high-level taxa in tropical America and Madagascar, and overlapping clades in South America, Africa, and Asia. The book concludes with a critique of adaptation by selection, founded on biogeography and recent work in genetics.
Jan 4, 2012

À propos de l'auteur

Michael Heads is a former Senior Lecturer in Ecology at the University of the South Pacific. He is now an independent scholar living in New Zealand.

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Molecular Panbiogeography of the Tropics - Michael Heads



The Species and Systematics series will investigate fundamental and practical aspects of systematics and taxonomy in a series of comprehensive volumes aimed at students and researchers in systematic biology and in the history and philosophy of biology. The book series will examine the role of descriptive taxonomy, its fusion with cyber-infrastructure, its future within biodiversity studies, and its importance as an empirical science. The philosophical consequences of classification, as well as its history, will be among the themes explored by this series, including systematic methods, empirical studies of taxonomic groups, the history of homology, and its significance in molecular systematics.

Editor in Chief: Malte C. Ebach (University of New South Wales, Australia)

Editorial Board

Sandra Carlson (University of California, Davis, USA)

Marcelo R. de Carvalho (University of Sao Paulo, Brazil)

Darren Curnoe (University of New South Wales, Australia)

Christina Flann (Netherlands Centre for Biodiversity Naturalis, The Netherlands)

Anthony C. Gill (University of Sydney, Australia)

Lynne R. Parenti (Smithsonian Institution, USA)

Olivier Rieppel (The Field Museum, Chicago, USA)

John S. Wilkins (University of Sydney, Australia)

Kipling Will (University of California, Berkeley, USA)

David M. Williams (The Natural History Museum, London, UK)

René Zaragüeta i Bagils (University of Paris 6, France)

University of California Press Editor: Charles R. Crumly



of the Tropics

Michael Heads


Berkeley • Los Angeles • London

University of California Press, one of the most distinguished university presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural sciences. Its activities are supported by the UC Press Foundation and by philanthropic contributions from individuals and institutions. For more information, visit www.ucpress.edu.

Species and Systematics, Vol. 4

For online version, see www.ucpress.edu.

University of California Press

Berkeley and Los Angeles, California

University of California Press, Ltd.

London, England

© 2012 by The Regents of the University of California

Library of Congress Cataloging-in-Publication Data

Heads, Michael J.

  Molecular panbiogeography of the tropics / Michael


    p.      cm.—(Species and systematics; v. 4)

Includes bibliographical references and index.

ISBN 978-0-520-27196-8 (cloth : alk. paper)

1. Biogeography—Tropics. 2. Biology—

Classification—Molecular aspects. 3. Variation

(Biology)—Tropics. I. Title.

QH84.5.H43 2012



19  18  17  16  15  14  13  12

10  9  8  7  6  5  4  3  2  1

The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 1997) (Permanence of Paper).∞

Cover photograph: Mist rising from the Borneo rainforest. Photo by Rhett A. Butler, mongabay.com.




1. Evolution in Space

2. Evolution in Time

3. Evolution and Biogeography of Primates: A New Model Based on Molecular Phylogenetics, Vicariance, and Plate Tectonics

4. Biogeography of New World Monkeys

5. Primates in Africa and Asia

6. Biogeography of the Central Pacific: Endemism, Vicariance, and Plate Tectonics

7. Biogeography of the Hawaiian Islands: The Global Context

8. Distribution within the Hawaiian Islands

9. Biogeography of Pantropical and Global Groups

10. Evolution in Space, Time, and Form: Beyond Centers of Origin, Dispersal, and Adaptation

Glossary of Geological Terms



About the Author


The theme of this book is the distribution of plants and animals and how it developed. The subject is approached using the methods of panbiogeography, a synthesis of plant geography, animal geography, and geology (Craw et al., 1999). The methodology is based on the idea that distribution is not due to chance dispersal; instead, range expansion (dispersal) and allopatric differentiation are both mediated by geological and climatic change. The biogeographic patterns discussed below mainly concern spatial variation in DNA, and so the subject can be termed molecular panbiogeography. The book focuses on molecular variation in plants and animals as this shows such clear geographic structure. Molecular analysis has revealed an intricate, orderly, geographic pattern in most groups examined, even in those that are apparently well dispersed, such as birds and marine taxa. This molecular/ geographic structure has often been described as surprising, as, for example, by Worth et al. (2010), reporting on bird-dispersed trees in Winteraceae, and it is certainly impressive. The discovery of this structure has been one of the most exciting developments in molecular biology, and it has intriguing, far-reaching implications for evolutionary studies in general. This book analyzes and integrates inherited information at the largest scale—the geographic distributions—and at the smallest scale—the molecular variation.

Molecular research has had a revolutionary impact on all aspects of biology and has led to revised ideas on the evolution and classification of many groups. Yet molecular variation is just morphology on a small scale, and there is no real conflict between the traditional morphological data and the new molecular data. Traditional taxonomic groups that were well supported in morphological studies are often corroborated in molecular work, and many of the radical realignments suggested by molecular studies are in groups and areas that morphologists have acknowledged as difficult. With respect to biogeography, most patterns shown in molecular clades were already documented in earlier systematic studies of some group or other.

In order to assess the reliability and importance of a proposed phylogeny, it is necessary to know many details of the particular study. These include the sample size, the part or parts of the genome sequenced, the methods of establishing sequences, the methods of analyzing them in order to produce a phylogeny, and the statistical support of the groups. These are not provided here because the book is not about these parameters. In the same way, accounts of biogeography using morphological taxonomy do not cite the morphological characters that were used to construct the taxonomies. The distributional and phylogenetic data cited herein are introduced as facts for discussion, that, hopefully, the reader will accept. This may not always be the case, but most of the studies referred to are exemplary accounts and most of the clades mentioned have good statistical support.

The first two chapters in this book deal with general aspects of interpreting evolution in space and time. The next eight chapters comprise a biogeographic transect around the tropics, from America to Africa, Asia, the Pacific, and back to America. The book does not give a systematic, area-by-area treatment, and only selected localities are covered in any detail. Australasia is covered in a separate volume. The main aim in this book is to provide worked examples and to illustrate principles using a new method of analysis. The groups that are discussed were chosen because their distributions are reasonably well known and they have been the subject of recent, detailed molecular study.


I am very grateful for the help and encouragement I've received from friends and colleagues, especially Lynne Parenti (Washington, D.C.), John Grehan (Buffalo), Isolda Luna-Vega and Juan Morrone (Mexico City), Jürg de Marmels (Maracay), Mauro Cavalcanti (Rio de Janeiro), Guilherme Ribeiro (Sao Paulo), Jorge Crisci (Buenos Aires), Andres Moreira-Muñoz (Santiago), Pierre Jolivet (Paris), Alan Myers (Cork), Robin Bruce and David Mabberley (London), Gareth Nelson and Pauline Ladiges (Melbourne), Malte Ebach (Sydney), Rhys Gardner (Auckland), Frank Climo and Karin Mahlfeld (Wellington), Bastow Wilson and Robin Craw (Dunedin), and Brian Patrick (Alexandra).


Evolution in Space

Many different ways of analyzing spatial variation in biological diversity—the biogeographic patterns—have been employed by different authors, and some of the assumptions in these methods are discussed here. The chronological aspect of evolution is discussed in the next chapter.

Every kind of plant or animal has its own particular distribution and ecology, and this was already well understood in ancient times. Yet portraying a distribution is not straightforward. New collections are always being made and ideas on the delimitation of taxonomic groups change. Outline maps are generalized simplifications only but are useful for comparative purposes. Although dot maps showing sample localities give more detail, they are always incomplete, the accuracy of the dot locations can often be questioned, and the entities that the dots represent—the populations or individuals—are constantly changing position due to birth, death, and movement. A distribution is dynamic and so a distribution map represents an approximation, a probability cloud, not an actual distribution. Nevertheless, the fact that so many distribution maps have been made reflects the high value that biologists and many others have put on them.

Knowledge of organic distribution is useful for simple survival and economic development, as the plants, animals, and microorganisms of a particular place are often among its most distinctive and valuable features, and also its most poisonous and dangerous. Many groups have particular, idiosyncratic distributions; the details of these are known by local people and broader-scale distributions are documented in the literature.

Organisms are distributed spatially in three dimensions and while the questions treated in this book mainly involve differentiation in the horizontal plane, in latitude and longitude, the altitudinal component of a clade's distribution must also be considered. While the elevation of a group is sometimes assumed to reflect its ecological preference, in some cases there is an ecological lag and historical effects are important. For example, an area may be uplifted along with its biota, and some of the biota will likely survive to become montane taxa. Depending on where it is located, a population may be uplifted or not during an episode of mountain building, and so biogeography can determine ecology, rather than the reverse.


The focus in this book is on distribution patterns and their interpretation in terms of evolutionary processes. Most biogeographic interpretation over the last 2,000 years has been based on a single paradigm, the center of origin/dispersal model of historical development. But having only a single working hypothesis to explain a set of phenomena can lead to problems, and over time it becomes easy to accept that the single hypothesis is the truth.

Although much modern work in biogeography stresses supposed consensus, in science and philosophy, as in art and literature, a diversity of views and approaches can be a good thing. Puritans of all sorts (whether Oliver Cromwell or Louis XIV) cannot stand anyone having a view different from their own. The inflexible schemes of these great simplifiers, levelers, and systematizers can hold up progress for decades. In contrast, geologists (Chamberlin, 1890, reprinted 1965) and now molecular biologists (Hickerson et al., 2010) cite the method of multiple working hypotheses, which proposes that it is never desirable to have just one working hypothesis to explain a given phenomenon. Accepting a single interpretation as definitive can be counterproductive and lead to the decline of a subject.

It is unfortunate that the interpretations of the data currently given in most molecular studies are all based on the same fundamental concepts. This plug-and-play biogeography involves the following steps: Assume that the study group has a center of origin and use a suitable program to find one; accept that fossil-calibrated clock dates give the maximum age of the group; describe possible dispersal routes from the center of origin. The axioms that are assumed here can be questioned, though, and a Socratic approach may be useful. Canetti (1935, reprinted 1962) wrote that A scholar's strength consists in concentrating all doubt onto his special subject, and a healthy scepticism is one of the pillars of science, both in history and in everyday practice. When identifying unfamiliar plants and animals on the reef or in the rainforest, it is tempting, but often dangerous, to jump to conclusions before considering a wide range of possibilities, and the same is true for biogeographic interpretation.

The case studies of different groups discussed below adopt certain assumptions and concepts, and some of these are outlined next.


A related group of organisms forms a branch or clade in a phylogeny or evolutionary family tree. A clade may or may not be be formally named as a taxon (plural: taxa). The closest relative of a group is termed its sister group. In most published phylogenies, the clades in a group are shown in a strictly hierarchical system of nested clades. Phylogeny is the general process of the evolution or genesis of clades, and a phylogeny is also a term for a branching diagram or a tree, a symbolic arrangement of hierarchical, nested sets of clades. Nested sets of groups are depicted in traditional dichotomous keys, nomenclatural systems, cladograms, phylogenies, trees, and so on; all represent the same thing, an Aristotelian classification. This is only one way of representing variation; another is ordination, a method which shows trends rather than groups and which is often used in ecology.

Ideas on the evolutionary process still reflect the Aristotelian, classificatory approach in many ways. This sometimes leads to a misplaced emphasis on the clades rather than on the morphological and molecular characters that underlie them. The usual units of analysis in this book are indeed clades, as presented in molecular phylogenies, but these should not be taken too literally. Biogeographic areas may be problematic, and biological groups—monophyletic clades—may also be complex. Most groups have characters/genes that show phylogenetic and geographic variation within the group that is incongruent with those of other characters/genes, and this will be discussed below. Ultimately, in a hypocladistic approach, the focus is on the evolution of the underlying characters rather than on particular combinations of characters, including the clades.


Evolution results in a continuum of differentiation. Entities may differ by a smaller or greater amount and, depending on this level, may be recognized as barely distinct populations, subspecies, species, genera, families, and so on. The focus here is on the process of differentiation rather than any of its particular products, and the species is seen here simply as a point on a trajectory between subspecies and genus; it has no special value. This is the species concept used by Darwin (1859) and Croizat (1964) (see also Ereshefsky, 2010). Neither the species nor any of the other taxonomic categories have any absolute value, and a species or genus in one group cannot necessarily be compared with species-or genus-level differentiation in another group.

In contrast with the Darwinian species, the species in the neo-Darwinian synthesis are very special indeed, as they have a reality that subspecies, genus, and the other categories do not. In this return to medieval nominalism, subspecies, genera, and other universals are seen as really just names and not things. Only species are real things (individuals). This distinction is not accepted in the Darwinian approach used here, in which clades (monophyletic groups) of any rank and their characters replace the species as the basic units of analysis. The most detailed information available on geographic differentation happens to concern monophyletic clades in morphological and molecular phylogenies, although geographic variation in any single character would be just as useful.


The interpretation of degree of difference (branch length) between groups is discussed in the next chapter. The particular degree of difference of a group and its taxonomic rank are not necessarily related to time; instead, they may reflect aspects of prior genome architecture in the ancestor. The focus here is on spatial differentiation in any clades, whatever rank or branch length is involved.

FIGURE 1-1. The distribution of hypothetical group A. Its center of origin might occur in the area with a highest diversity in the group (1), in the region of the oldest fossil (2), in the area of the most advanced form (3), in the area of the most primitive form (4), or in the area of the basal group (5).


The center of origin/dispersal model and the vicariance model are often contrasted in biogeographic studies, and their applications have caused a great deal of debate.

Center of Origin/Dispersal Model

How did the distribution of a plant or animal develop? Consider a hypothetical distribution (Fig. 1-1). One theory is that such a pattern originated by a plant or animal evolving at a point somewhere within its current area and spreading out from there to the limits of its present range. Researchers attempt to locate the center of origin or ancestral area by studying the distribution and phylogeny in the group itself and by using different criteria. The center of origin has been thought to occur in the area that shows one or more of the following:

the highest diversity of forms within the group (1 in Fig. 1-1),

the oldest fossil (2 in Fig. 1-1),

the most advanced form (cf. Darwin, 1859; Briggs, 2003) (3 in Fig. 1-1),

the most primitive form (cf. Mayr, 1942; Hennig, 1966) (4 in Fig. 1-1), and

the basal clade or grade of the group (most modern studies) (5 in Fig. 1-1).

Several computer programs designed to find the center of origin of a group are now available, for example, DIVA (Ronquist, 1997) and Lagrange (Ree and Smith, 2008).

Many other criteria for locating a group's center of origin have been proposed in addition to those listed above, and the confusion that this implies was pointed out by Cain (1943). This paper led to the modern critique of the center of origin that has been developed in panbiogeography (Craw et al., 1999) and paleontology (López-Martínez, 2003, 2009; Cecca, 2008).

Cecca (2008) characterized two models of evolutionary biogeography: center of origin/dispersal theory as developed by Darwin (1859) and Wallace (1876), and vicariance, as developed by Sclater (1864) and Croizat (1964). In discussions of these models, the phrase center of origin does not simply mean a center where a group has originated (all taxa originate somewhere), but refers to a specific concept used in the dispersal model. In this model, a group's ancestor evolves as a monomorphic, homogeneous entity in a restricted area (the center of origin) following a chance dispersal event, and the group attains its distribution by physical movement out of this center.

Vicariance Model

Finding the center of origin of a group is a fundamental aim of many studies, and groups may be analyzed in ever-increasing detail in order to locate the center. The center of origin of a group is often located by examining the group itself. An alternative approach considers a group not on its own, but in relationship to its closest relative or sister group (Fig. 1-2). It may be difficult to understand the origin of a group by studying the group itself, especially if groups come into existence together with at least one other, by vicariance.

In many cases two sister groups have neatly allopatric (vicariant) distributions, with one group replacing or representing the other in a second area, often nearby or even adjacent to the first. Each of the two groups may have arisen not by spreading out from a point, but by geographic (allopatric) differentiation in its respective area from a widespread ancestor. In this process (vicariance), there is no physical movement, only differentiation, with populations in area A evolving into one form and populations in area B into another. This vicariance theory is a basic component of panbiogeography. In a vicariance event, the distribution of a group comes into existence with the group itself. A group's center of origin may be more or less the same as its distribution, especially if it is part of an allopatric series. (A distribution range may expand or contract after its initial formation, leading to secondary overlap; this is discussed below.)

FIGURE 1-2. Group A and its sister group, B, two allopatric clades.

Despite the development of the vicariance model, the center of origin/dispersal model of the evolutionary process is still widely assumed by paleontologists (Eldredge et al., 2005), ecologists (Levin, 2000; Gaston, 2003: 81), and some biogeographers. For example, Cox and Moore (2010: 204) wrote: "Let us imagine that a species has recently evolved. It is likely, to begin with, to expand its area of distribution or range until it meets barriers of one kind or another." But this does not necessarily happen in a vicariance event, as the new species already abut their relatives. If a globally widespread form evolves by breaking down into, say, two allopatric species, one in the northern hemisphere and one in the southern hemisphere, neither one may expand its range. In a vicariance model, a new clade is just as likely to contract its range as to expand it.

In the vicariance approach, the focus is on tracing the originary breaks between groups, not on locating a point center of origin within a group. In a dispersal analysis, the first question is: Where is the center of origin? In a vicariance analysis, the first question is: Where is the sister group? The focus is not on the group itself or on details of its internal geographic/phylogenetic structure, but on its geographic and ecological relationship with its sister group and other relatives.

In this model, a group originates by the breakdown of a widespread ancestor, not by evolving at a point and spreading out from there. Analysis of any group can start either with a point center of origin or, alternatively, with a widespread ancestor. In the latter model, a group evolves on a broad front over the region it occupies, by fracturing with its sisters (vicariance) at phylogenetic and biogeographic breaks or nodes. A node is not a center of origin or an ancestor; it is a break where the distributions of two or more groups meet.

In one example, Chakrabarty (2004) supported a vicariance history for the freshwater fish family Cichlidae. He compared the process with a mirror being struck several times with a hammer. There is no movement of the individual shards, which are all neatly vicariant. The sequence of the hammer blows (i.e., phases of differentiation) is seen in the phylogeny or cladogram. The process also resembles the development of vascular tissue in a young organ out of ground tissue. There is no physical movement and the veins do not grow by pushing their way through tissue, but by differentiating in situ, in accordance with the genetic program.

Modes of Speciation: Dispersal and Vicariance

The mode of differentiation of groups in general (phylogenesis) and of species in particular (speciation) is problematic, and the interpretation of even the simplest cases is debated. As with clades in general, two allopatric species can be explained as the result of either vicariance in a widespread ancestor (dichopatric speciation) or founder dispersal from a center of origin (peripatric speciation).

Origin of the Ancestor

Two descendant groups may have originated by vicariance, but what about the ancestor of the two? Surely the ancestor must have dispersed to achieve its wide range? In fact, this is not necessary, as the ancestor of the two groups A and B, in areas A and B, may itself have originated as an allopatric member of a broader complex, the ancestor of A + B + C, that also occurred in area C. This in turn may have differentiated from the ancestor of A + B + C + D, as indicated by the four clades in Figure 1-3. Here there is no center of origin. In the center of origin/dispersal theory, each of the four allopatric groups in Figure 1-3 would have a separate center of origin, and their distributions are not directly related to their origins—the groups formed first and the distributions were established later. The boundaries of the four groups are secondary and the distributions only met after the four groups spread out from their respective centers of origin. Instead, in panbiogeography the mutual boundaries are interpreted as phylogenetic and geographic breaks or nodes. These recur at the same localities in many different groups with different ecology, and so a chance explanation is unlikely.

FIGURE 1-3. Four groups A-D with allopatric distributions in areas A-D. The phylogeny of the four groups is also shown.

Overlap in Distribution

In many cases, the allopatry between close relatives is not perfect and the groups show marginal overlap or interdigitation. This can represent local, secondary overlap by range expansion following the original, allopatric differentiation.

Sometimes a group shows extensive secondary overlap with its sister; this occurs mainly at higher taxonomic levels such as families and orders. A worldwide group and its worldwide sister may have each occupied half the Earth before the overlap developed. A vicariance analysis of the groups' history does not involve finding each group's center of origin, but tracing possible original breaks between the two groups. For example, one of the worldwide sister groups may be fundamentally northern, the other southern.

A few species, more genera, and many higher-level taxa, such as birds and flowering plants, are worldwide and overlap with each other everywhere. This pattern probably reflects the older age of the higher categories. The overlap of groups shows that vicariance cannot be the only biogeographic process. If it were, there would be pure allopatry—each point on Earth would only have one, locally or regionally endemic, life form. As it is, most places have biotas that include many kinds of plants and animals, indicating the overlap of clades. The overlap may be due to early phases of large-scale range expansion by whole communities.

If the affinities of any group are traced far enough, these will be found to make up a worldwide complex. Beyond this stage, if not before, there must be overlap with relatives. The widespread overlap among, for example, families and orders of birds indicates phases of range expansion. In primates, for example, the main branches—Old World monkeys, New World monkeys, lemurs, tarsiers—are notable for their high degree of allopatry. On the other hand, primates as a whole (together with their close relatives) show wide overlap with their sister group, the rodents and lagomorphs. At some stage (probably between the origin of primates and the origin of the main primate subgroups) there was a phase of overlap between the proto-primate complex and the proto-rodent complex.

Phases of vicariance affecting whole communities are often attributed to geological changes in the past, such as the opening of the Atlantic. In the same way, phases of population mobilism, with range expansion and colonization, probably occurred during and following the great geological revolutions. For example, the last, great phase of marine transgressions, in the Mesozoic, produced dramatically extended coastlines and associated habitats. Marine transgressions occurred on all the continents and, at the same time, rifting and continental breakup also produced new seaways. During this phase of mobilism, groups with suitable coastal, marginal ecology colonized vast areas of new habitat and became widespread globally. This was followed by a phase of immobilism through the Cenozoic, during which local differentiation predominated.


It is often suggested that the main factor distinguishing evolution by vicariance and by dispersal is the time of appearance of the barrier—before evolution in dispersal and during evolution in vicariance. A more general point is that in a vicariance model, the Earth and its life evolve together, whereas in the dispersal model they do not. In dispersal theory, every taxon has its own unique history caused by one-off, chance events, and there are no community-wide biogeographic patterns with single causes. (Molecular clock studies based on dispersal theory also support this notion; this will be discussed in Chapter 2.) Yet many biologists would be reluctant to abandon the idea that geology can cause community-wide vicariance and generate both large-and small-scale community patterns. The idea that Earth and life evolve together is seen in the geographic concordance of many aspects of biogeography. This agreement even occurs among groups with completely different ecology, such as intertidal marine groups and montane groups. To cite just one example, the Hawaiian Islands and the Marquesas Islands form a center of endemism that is unexpected, given the direction of the currents, and yet it is defined by reef fishes, insects, and montane plants (see Chapter 7). Geographic congruence among different groups and also a general congruence between biogeography and geology have been known for a long time. For example, The primary geographical divisions in the global mammal fauna clearly coincide with geology and plate boundaries (Kreft and Jetz, 2010: 19). For example, in mammals, major phylogenetic/geographic breaks (nodes) occur between South America and Africa, and between Madagascar and Africa.


Four key processes have been proposed in biogeography. As discussed above, differentiation (e.g., speciation) can be due to vicariance of a widespread ancestor or to founder dispersal from a center of origin. In addition, two overlapping sister clades can be explained as the result of range expansion (by normal ecological dispersal, simple physical movement) or by sympatric differentiation.

Of the four processes just cited, vicariance and normal ecological dispersal are accepted as important by all authors. They are the two processes that are accepted in this book as explaining distributions. Normal ecological dispersal can involve movement within the distribution area or outside it, and this may lead to range expansion. Range expansion explains overlap; it does not explain allopatry.

The third process, sympatric differentiation, was controversial, although it is now accepted in some cases (Schluter, 2001; Friesen et al., 2007; Bolnick and Fitzpatrick, 2007). If it does occur, it is probably quite rare and many cases of supposed sympatry between sister groups prove, on closer examination, to involve only partial geographic overlap and significant allopatry. Other apparent cases of sympatry may involve allopatry at a small scale. As noted above, low-level clades are often allopatric with their sisters, whereas higher-level clades show more overlap, so overlap can generally be regarded as a secondary process that has developed over time from original allopatry, rather than by sympatric evolution.

The fourth process, differentiation by founder dispersal, is controversial and may not exist.


Three quite different processes have all been termed dispersal, and they can be contrasted as follows.

Normal Ecological Dispersal. This is the normal physical movement seen in plants and animals. It includes daily and annual migrations, along with the dispersal of juveniles. The movement is made possible by the well-known mechanisms observed in different groups. Normal ecological dispersal occurs every day and does not lead to differentiation (speciation, etc.). It may take place over long distances—for example, in sea-birds—or over much shorter distances, depending on the organism. Following their origin by reproduction, all individual organisms have dispersed to where they are by this process. Normal ecological dispersal is seen in the weeds that soon colonize a disturbed area, whether this is a newly dug garden, an area of burnt vegetation, the area in front of a retreating glacier, a landslide, or a volcanic island such as Krakatau in Indonesia that has been devastated by a recent explosive eruption.

Despite appearances, this process of simple movement does not necessarily explain the distribution area occupied by a taxon—in particular, any allopatry with related taxa—as it does not account for evolutionary differentiation, and this can, by itself, produce a distribution. Thornton and New (2007) titled their book Island Colonization: The Origin and Development of Island Communities, yet studies on the colonization of Krakatau, for example, only concern the ecological origin and development of communities, not their evolutionary origin. The community on Krakatau is a subset of the weedy community that already existed on the islands in the region, and its evolutionary origin dates to long before the last eruption on Krakatau.

Range Expansion. Following a normal dispersal event, an organism's new position may lie within the former range of the taxon or it may lie outside it and represent a range expansion. Range expansion is seen in historical times in the anthropogenic spread of weeds and at other times in geological and evolutionary history. Range expansion, when it does occur, may be very rapid and a more or less local plant or animal may become worldwide in hundreds rather than millions of years. This takes place by normal ecological dispersal using the normal means of dispersal in the group, not the rarely used or unknown means sometimes cited to explain the more spectacular events of founder dispersal.

A global ancestor may have achieved its range during Mesozoic range expansion. This mobilism eventually stabilized and was replaced with a phase of immobilism through the Cenozoic. This was a period of in situ evolution that produced local differentiation, mainly at species and subspecies levels. Phases of mobilism may alternate with phases of immobilism in which allopatric evolution (vicariance) takes place. Earlier phases of population mobilism would have occurred as new landscapes emerged from the devastation of the Permo-Carboniferous ice ages, centered in the southern hemisphere and much more severe and long-lasting than the Pleistocene ice ages. These cycles of biogeographic mobilism and immobilism may take tens of millions of years to complete, as with the geological cycles of mountain uplift, erosion, deposition, and further uplift.

Naturally, it is far more difficult to analyze the biogeography of a time prior to the one in which the modern, extant patterns developed, and many aspects of the premodern patterns will never be known. The modern centers of endemism, in their turn, will not last forever. A new geological or major climatic catastrophe will eventually lead to massive extinction and renewed mobilism, with weedy taxa taking over before they settle and establish new regional blocks of endemic taxa.

Long-distance Dispersal/Speciation by Founder Dispersal. The defining feature of this process is not so much the long distance but the fact that it involves a unique, extraordinary dispersal event by a founder across a barrier. This leads to isolation and speciation (or at least some differentiation). As Clark et al. (2008) emphasized, there is an important distinction between dispersal as normal individual movement and range expansion on one hand, processes that are seen every day, and long-distance dispersal involving founder speciation on the other. The latter (termed dispersal-mediated allopatry in Clark et al., 2008) is a theoretical construction. Normal ecological movement and range expansion, along with other kinds of dispersal such as daily and annual migrations, are accepted here; long-distance dispersal/founder speciation is not.

As noted, every individual plant and animal moves as part of its normal means of survival, at least during one stage of its life cycle. With the exception of some colonial taxa, all individual organisms have reached their present position by dispersing there. This normal ecological movement should not be confused with long-distance or founder dispersal, which leads to new lineages. Dispersalists argue that When lineages arrive in new habitats they will usually diverge and sometimes speciate (Renner, 2005). But any patch of newly cleared garden will soon be colonized by weedy flora and fauna, later by less weedy taxa, and none of these will speciate there. Again, founder dispersal is quite distinct from normal dispersal. Authors supporting the center of origin/ founder dispersal view for one or other group have often concentrated on proving that ecological dispersal, or ordinary movement, does occur, but this may not be relevant to the issue of founder dispersal.

For differentiation or speciation to occur, a fundamental change in the population ecology from a state of mobilism to one of relative immobilism has to occur. Dispersal on its own might explain how primates came to be in America if the American primates were the same as those of Africa or Asia. But physical movement on its own cannot explain why the American primates are different and form their own group. One main problem with founder dispersal is explaining how movement between populations could be occurring at one time, but then at some point stop or at least decrease (leading to differentiation). What is the reason for the crucial change from high rates of dispersal to low rates? In theory, this might be due to changing behavioral patterns in animals or means of dispersal in plants, but this cannot explain repeated patterns in unrelated animals and plants. Geological or climatic change is one obvious possibility, and this is the basis of vicariance. In center of origin theory, dispersal and speciation are instead determined by chance—the change from movement to no (or less) movement is created by a barrier which is permeable to a chance crossing by a single founder, but is then, somehow, impermeable to all others. In this view, the evolutionary biogeography of a group is due to chance, and so there is no need to examine any details of distributions that cannot be attributed to local ecology.

Dispersalists have sometimes suggested that island endemic taxa, for example, had much more effective means of dispersal in the past than they do now, and that these were lost with evolution, trapping taxa on an island (Carlquist, 1966a, 1966b). This is an ingenious and logical solution to the general problem that is often not mentioned—what causes the change from a phase of dispersal to a phase of no dispersal? Unfortunately, the idea of loss of means is probably wrong, as most endemics in most places have not lost their means of dispersal. But the fact that the idea was proposed at all indicates there is a problem that cannot be solved simply by citing chance. Profound geological and ecological change is a more likely reason for cycles of immobilism-mobilism-immobilism.

Dispersal theory accepts that normal ecological dispersal and founder dispersal both occur in nature, whereas vicariance theory only accepts the first, but in any case it is important to distinguish between the two processes. The fact that a distinction is not made between contiguous range expansion (by normal dispersal) and across-barrier, founder dispersal is a serious drawback with programs such as DIVA (Kodandaramaiah, 2010). This conflation of the two different processes is a defining feature of dispersal biogeography (Matthew, 1915) and is also the basis of the confusing criticism that panbiogeography denies dispersal.

To summarize: Most modern biogeographers follow Mayr (1982, 1997) in accepting that allopatry can be caused either by vicariance (dichopatry) or by founder dispersal (peripatry), but only vicariance is accepted here. Allopatry is accounted for by immobilism and vicariance, while overlap among groups can be attributed to range expansion and population mobilism (‘dispersal’).

Dispersal: Any and All Changes in Position

Many birds, primates, and other groups show daily and annual migrations that involve significant distances and are repeated through the millennia. These need to be accounted for in biogeography and ecology, although they are usually dealt with separately. Clements and Shelford (1939) realized the problem and introduced the highly generalized, rigorously geometric concept of any and all changes in position. This would include changes in position due to physical movement or to evolution. The authors' suggestion that this concept be termed dispersal or migration was elegant but confusing and never caught on. This does not detract from the value of the concept. As with global phylogenies and the evolution of major groups, daily migrations of animals, even at a local scale, may reflect either current ecological conditions or past features such as former streams, rivers, or coastlines.


A phylogeny often has its main division, its basal break, between a small group and a more diverse sister group containing several clades. The smaller group is termed basal, although strictly speaking only the nodes or breaks between groups are basal; no group is more or less basal than its sister group. The term basal group is thus potentially misleading, as a basal group is no more primitive than its sister, and is not ancestral to it (Krell and Cranston, 2004; Crisp and Cook, 2005; Santos, 2007; Omland et al., 2008). Nevertheless, by now the term basal group is widely used and understood in its purely topological sense, and it is a useful term for a smaller sister group. It should probably always be used in quote marks, to indicate the problem, but then terms such as clade, monophyletic, dispersal, center, gene, and so on would have to be treated in the same way.

The phrases sister to the rest of and basal in are used here more or less interchangeably. The difference between the two is arbitrary and mainly nomenclatural—a basal group is considered to be part of the sister group and has the same name; a sister is a separate group and has a different name.

Although an ancestor would be basal in a phylogeny, a basal group is not necessarily ancestral or structurally primitive. For example, Amborella is likely to be the basal angiosperm, sister to all the rest, but Pennisi (2009: 28) went one step further and suggested: "Given that placement, Amborella's tiny flowers may hint at what early blossoms were like." In fact, there is no reason why one (Amborella) or the other (all the other flowering plants) of the two sister branches should have a flower that is more primitive.

In the same way, the basal clade in a group is often interpreted as occupying the center of origin for the group, although this cannot be justified (Crisp and Cook, 2005). Likewise, morphological analysis may show that the oldest fossil clade in a group is phylogenetically basal to the rest, but it cannot be assumed to be ancestral to the others; it may simply be an extinct sister group.

Thus the idea that basal groups in a phylogeny are ancestral can be rejected as a generalization. Basal groups are simply less diverse sister groups, and their distribution boundaries may represent centers of differentiation in what were already widespread ancestors, not centers of origin for the whole group (Heads, 2009a).

Despite these arguments, modern phylogeographic studies often assume that a basal clade is primitive, ancestral, and located near the group's original center of origin, while advanced members of a clade have migrated away (Avise, 2000). This idea is derived from Mayr (1942) and Hennig (1966), who proposed that the primitive member of a group occurs at the center of origin. This is in contrast with the Darwinian model, which assumes that an advanced form would outcompete the older forms and force them to migrate away (Darwin, 1859; Matthew, 1915; Darlington, 1966; Frey, 1993; Briggs, 2003). In this view, the center of origin is occupied by derived forms. The conflict between the Darwinians and the Mayr/Hennig/phylogeography school over the center of origin is irrelevant in the vicariance model, where there is no center of origin to begin with.

To summarize, a basal group is not ancestral; it is simply the smaller of two sister groups. Both will have the same age and neither one is derived from the other, or more advanced or primitive than the other.


Good examples of basal group/center of origin analyses are seen in the extensive literature proposing dispersal into and out of the Caribbean. In the mockingbirds, Mimidae, a Yucatán endemic is basal to a largely Caribbean clade. Lovette and Rubenstein (2007: 1045) argued that this is suggestive of a pathway of colonization into the Antilles from central America via Cuba. Conversely, in butterflies, the Greater Antilles genus Antillea is basal to a widespread clade (Phyciodina) of North, South, and Central America. So from a center of origin in the Antilles, The ancestral Phyciodina colonized the [Antilles-Venezuela] landspan and spread south to the Guyanan Shield and then quickly to the Brazilian Shield (Wahlberg and Freitas, 2007: 1265). (The subsequent scenario involved a convoluted history of transcontinental dispersals and back-dispersals, although the authors described these butterflies as well-known to be relatively sedentary.)

In fact, no migration into or out of the Caribbean is required for the mockingbirds or the butterflies. In both groups, the location of the basal clade in the Yucatán/Greater Antilles region and the distribution of the rest of the group elsewhere can be explained by simple vicariance somewhere around Yucatán/Greater Antilles in an already widespread ancestor. The basal node represents an early center of differentiation in already widespread groups, not a center of origin. In a similar example, Sturge et al. (2009) wrote that molecular phylogeny confirms that the New World oriole Icterus (Icteridae) colonized South America from the Antilles, but this was only because South American species were nested in an otherwise Antillean clade and a widespread ancestor is a more parsimonious solution.


Consider the group of four taxa A–D shown in Figure 1-3 that are found in allopatric areas A–D and have a phylogeny (D (C (B + A))). In modern dispersal theory, the sequence of nodes in a phylogeny is read as a sequence of dispersal events, with taxa invading a new region, differentiating there, and then invading another region. The center of origin is occupied by the basal population in the basal group, D, and the phylogenetic sequence reflects a series of dispersal events from area D to C, B, and A. Each of the four taxa has its own individual center of origin somewhere within its range.

The model has been criticized by vicariance biogeographers because the simple allopatry among the four clades might not be due to dispersal but to a sequence of in situ differentiation events in an ancestor that was already widespread in A–D. The phylogeny (D (C (B + A))) would then reflect a sequence of breaks among the areas: D versus A + B + C, C versus A + B, A versus B. In this case, the sequence of differentiation shows a simple progression from east to west in Figure 1-3, and there is no physical movement.

In other cases, the phylogeny does not follow a simple geographic progression. Figure 1-4 shows a pattern in which the two sequential basal clades in a group, A and B, are not adjacent geographically and are separated by other groups, C and D. Dispersal theory would attribute this to jump dispersal. In vicariance theory it indicates that a widespread ancestor differentiated first at two basal nodes (between A and the rest, then between B and C + D) and finally at a node geographically between A and B.

Thus a phylogeny may convey the impression of a center of origin at the locality of the basal clade and dispersal from there, but if there was a widespread ancestor this is not necessary. Major disjunctions of tens of thousands of kilometers often occur between taxa at consecutive nodes on a phylogeny, even in groups in which long-distance, colonizing dispersal is improbable. Here it is especially likely that a phylogeny reflects a sequence of vicariance events. In many groups, differentiation has taken place repeatedly and more or less simultaneously around just a few globally significant nodes, such as the southwest Pacific basin and southwest Indian Ocean basin. Consider a phylogeny in five groups: Australia (Madagascar (Australia (Madagascar (Australia)))). In a dispersal interpretation, this would require repeated long-distance dispersal events backward and forward between the two centers. A vicariance interpretation of the same pattern proposes repeated differentiation at the same nodes in a widespread ancestor, perhaps caused by reactivation of tectonic features.

FIGURE 1-4. Four groups A-D with allopatric distributions in areas A-D. The phylogeny of the four groups is also shown.

Vicariance interpretations of phylogenetic sequences are given in some recent literature. For example, in the New World snake Bothriechis, earlier studies deduced a process of dispersal from Costa Rica to southern Mexico. Instead, Castoe et al. (2009: 98) interpreted the phylogeny as indicating "a more simplistic northward progression of cladogenesis that requires no inference of dispersal (italics added). A similar pattern is also seen in other snakes of the area, suggesting vicariance as the primary driving force underlying speciation." The Great American Biotic Interchange theory proposes dispersal across Central America, so the idea that evolution in the region may not have involved physical movement is of special interest. In another example, Doan (2003) interpreted a phylogeny of Andean lizards as reflecting a northward sequence of speciation in a widespread ancestor, rather than northward dispersal. In a botanical study, the phylogeny of Rhododendron (Ericaceae) in Malesia was interpreted as a geographical progression of cladogenesis (Brown et al., 2006). The same method of interpreting phylogeny used in these papers is adopted here.

Summing up, phylogenies of extant clades can indicate a sequence of divisions (nodes) between sister groups, rather than a sequence of ancestors and descendants. Dispersalists have adopted the second option, but this has only led to long-lasting, unresolved debates about the center of origin in particular groups, about how to locate the center of origin in the first place, and what the means of dispersal could be. In the model of evolution proposed here, there is no center of origin (other than the point of break, which is a margin rather than a center), there is no founder dispersal speciation, and there is no radiation from a center. If the ancestor is already widespread geographically (and probably also ecologically) before the differentiation of the descendant groups, the issue is no longer about how the modern groups reached a certain area, but how they evolved there—in other words, where the breaks occurred that led to their differentiation. Once the spatial context is clarified, the question of timing should be more straightforward.


In the dispersal-vicariance analysis of Ronquist (1997), inferences of dispersal events are minimized as they attract a cost. Extinction also attracts a cost, but vicariance does not. It was not explained why this approach should be taken and, as suggested above, it is based on a confusion of the two different concepts of dispersal. Dispersal in the sense of ordinary movement should not attract any cost in any model. Jump or founder dispersal would attract no cost in a traditional dispersalist model, although in a vicariance model of speciation or evolution it is rejected a priori.

In most modern studies, the spatial analysis of phylogeny has been based on the idea of a center of origin, and so authors employ programs, such as DIVA, that will often find one. Authors looking for a particular center of origin sometimes complain that DIVA will find a widespread ancestor if, for example, all the extant groups are allopatric. But even when they are not, a widespread ancestor can still be proposed, as original allopatry may have been obscured by subsequent range expansion or extinction. There is no logical need to interpret a phylogeny as a series of dispersal events.


If one group occurs on a mainland, a, and its sister group occurs on a much smaller island, b, (Fig. 1-5), the island group is often assumed to have been derived from the mainland group by dispersal. The island forms are predicted to be related to particular populations in their large sister group. Yet well-sampled molecular studies now show that in many of these cases the phylogeny has the pattern: (a¹, a², a³…) (b¹, b², b³…), where the superscripts indicate different areas within a and b. The groups in the two areas are reciprocally monophyletic and the group in b is not related to any one population in a. In this type of pattern, dispersal can still be salvaged as an explanation, but only if it occurred prior to any other differentiation in the groups, and this is often unlikely. On the other hand, reciprocal monophyly is the standard signature of simple vicariance of a widespread ancestor at a break between a and b. Even if groups in the two areas a and b are not reciprocally monophyletic, vicariance is still possible, and this is discussed next.

FIGURE 1-5. Two sister groups, a and b, on a mainland and an island. The phylogeny is: (a¹ + a² +…a⁷) (b¹ + b²), and the mainland clade and the island clade are reciprocally monophyletic.


A monophyletic clade includes all the branches derived from a single node. A paraphyletic group or grade comprises several sequential branches of a phylogeny, but does not include all the branches derived from a node (e.g., in Fig. 1-3, the clades B, C, and D, but not A). Many groups comprise a basal grade located in one area, A, and a disjunct population or clade in a second area, B. The pattern is usually explained as the result of dispersal of the clade from A to B. Instead, a grade located in a single area may represent a phase of differentiation there, not a center of origin (Heads, 2009b). For example, within a widespread ancestor already in A and B (Fig. 1-6), allopatric evolution may occur around a node at A. If this is followed by secondary overlap at A and extinction of populations between A and B, this will produce a basal grade in area A (Fig. 1-6D). In actual cases, the overlapping clades in area A often show slight but significant differences in their distribution (as indicated in Fig. 1-6D), and these may represent traces of the earlier phase of allopatry. Many biogeographic analyses treat all the species in an area such as A as having the same distribution, overlooking any allopatry, and this can confuse analysis. To summarize: A dispersal analysis interprets the pattern described here, with a basal grade in A, by long-distance dispersal from A to B across a barrier. Instead, a vicariance analysis infers differentiation in a widespread ancestor followed by local overlap within A by normal means of dispersal.

FIGURE 1-6. A hypothetical example of distribution in a taxon currently found in two areas. A. A widespread ancestor. B. This begins to differentiate around a node (star) associated with the formation of a mountain range or inland sea, for example. C. The ancestor has differentiated into five allopatric clades, four with a narrow range and one widespread. Their ranges begin to overlap while some of the populations of the widespread clade suffer extinction (broken line). D. The clades now overlap but the ranges still show traces of their original allopatry. Following extinction of populations between areas A and B, the outlier in B may appear to be a secondary feature and the result of long-distance dispersal.

FIGURE 1-7. The phylogeny and distribution of clades in the Arctotidinae (from Funk et al., 2007).

Examples of Groups with Basal Grades in One Area

The Arctotidinae (Asteraceae) are a good example of the basal grade pattern. The group comprises a basal grade of three southern African clades as well as Cymbonotus of southern Australia, which is sister to two other southern African genera (Fig. 1-7; Funk et al., 2007). Cymbonotus is sister to a southern African clade and embedded in an Australian-southern African clade; it is not embedded in a southern African clade. It is likely that the southern African clades show significant differences in their distributions within the region.

In another example, Protea (Proteaceae) is diverse in the Cape region of South Africa and also has a few species widespread throughout tropical Africa. Valente et al. (2010) found that non-Cape species are nested within a wider radiation of Cape lineages and all except two of them belong to a single clade. Therefore, the authors suggested, most extant lineages outside the Cape originated by in situ diversification from a single ancestor that arrived there from the Cape (p. 746). This logic is not accepted here (cf. Fig. 1-6). The authors only interpreted the phylogeny in terms of a dispersal model because of the programs they used; an alternative vicariance scenario was not considered. Nevertheless, the sister genus of Protea is Faurea, centered in tropical Africa (where Protea has low diversity) and Madagascar (where Protea is absent); a vicariance analysis would be simple and of considerable interest.

In a third example, Grandcolas et al. (2008: 3311) argued that within certain New Caledonian groups, multiple species are nested within larger clades with taxa from Australia, New Zealand or New Guinea, calling for explanations in terms of recent dispersal [to New Caledonia]. Thus the phylogeny: (Australia (Australia (Australia (New Caledonia)))) is taken to reflect a center of origin in Australia, where the basal grade occurs. An alternative explanation for the pattern would be differentiation in a widespread Australian-New Caledonian ancestor, as in Figure 1-6.

Center of Origin/Basal Grade Theory and Ancestral Habitat Reconstruction

As discussed, a basal grade does not necessarily indicate a geographic center of origin, and this argument also applies to ecology, as a basal grade is often thought to occupy the ancestral habitat. For centropagid crustaceans, Adamowicz et al. (2010) concluded that Species occupying saline lakes are nested within freshwater clades, indicating invasion of these habitats via fresh waters rather than directly from the ocean or from epicontinental seas (p. 418). But, using the same argument given above for geography, this ecological phylogeny: (freshwater (freshwater (freshwater (saline lakes)))), does not necessarily mean that freshwater habitat was the center of origin for the saline lake clade. The ancestor of the whole group may have occupied both freshwater and saline lakes and itself be derived from a marine ancestor, for example, following marine transgression and regression.

In other crustaceans, the spiny lobsters, Palinuridae, are widespread in warmer seas and especially common around Australasia. Tsang et al. (2009) reasoned that the three genera restricted to the southern high latitudes (Jasus, Projasus, and Sagmariasus) are the basal lineages in the family, suggesting a Southern Hemisphere origin for the group. In the same way, the authors assumed that the basal groups indicated the ecological center of origin. For one clade, they wrote, "the shallow-water genus Panulirus is the basal taxon in Stridentes, while the deep-sea genera Puerulus and Linuparus are found to be derived. This indicates that the spiny lobsters invaded deep-sea habitats from the shallower water rocky reefs and then radiated." Again, the habitat of the basal taxon is not necessarily a center of origin for the other groups; the ancestor may have already been widespread in both deep and shallow water before it differentiated into the modern genera.


Many groups have a basal grade in one region, as in the last pattern, and also have a widespread distal clade.


The dandelion tribe Cichorieae (= Lactuceae) of Asteraceae is cosmopolitan and has its basal clades and also its sister group (Gundelieae), that is, a basal grade, in the Mediterranean region (Funk et al., 2005). As usual, this does not mean the Mediterranean basin was a center of origin from which the tribe itself spread. Instead, the region may have been an early center of differentiation in an already global ancestral Cichorieae.


The family Asteraceae as a whole, the largest plant family, with ~24,000 species, is another example of a group with a basal grade in one region and a widespread distal clade. Its basal group, subfam. Barnadesioideae, is in South America. The sister groups of the family are centered on the eastern Pacific margin (Calyceraceae of South America) and western Pacific (Goodeniaceae, mainly in Australia;

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